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Development of a Rain Down Technique to Artificially Infest Hemlocks with the Hemlock Woolly Adelgid, Adelges Tsugae Abstract

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Development of a Rain Down Technique to Artificially Infest Hemlocks with the Hemlock Woolly Adelgid, Adelges Tsugae Abstract Journal of Insect Science: Vol. 14 | Article 106 Jetton et al. Development of a rain down technique to artificially infest hemlocks with the hemlock woolly adelgid, Adelges tsugae Robert M. Jetton1a, Albert E. Mayfield III2b, and Zaidee L. Powers3c 1Research Assistant Professor, Camcore, Department of Forestry & Environmental Resources, North Carolina State University, Raleigh 2Research Entomologist, USDA Forest Service, Southern Research Station, Asheville, NC 3Graduate Research Assistant, Camcore Department of Forestry & Environmental Resources, North Carolina State University, Raleigh Abstract The hemlock woolly adelgid Adelges tsugae Annand (Hemiptera: Adelgidae), is a non-native invasive pest that has caused widespread decline and mortality of eastern hemlock (Tsuga canadensis (L.) Carr. (Pinales: Pinaceae)) and Carolina hemlock (T. caroliniana Engelm.) in the eastern United States. Our pre- liminary experiments evaluated the utility of a rain-down technique to induce artificial infestations of A. tsugae on hemlock seedlings en masse. Experiments were conducted in PVC (1 m3) cages topped with poultry wire for placement of A. tsugae-infested branches, and with 1 m2 gridded glue sheets and/or hem- lock seedlings placed below to capture adelgid abundance, distribution, and infestation rate data. In the March 2011 experiment, the density of progrediens crawlers (adelgid nymphs, first instars) that rained down inside the PVC cages was significantly higher in the high ovisac treatment compared to the low ovi- sac treatment, with an estimated 513,000 and 289,000 crawlers per m2 falling beneath each treatment, respectively. Resulting A. tsugae infestation rates on Carolina hemlock seedlings placed inside the cages did not differ between the treatments but were at or above established damage threshold densities for the adelgid. Infestation rates on eastern hemlock seedlings that were placed in cages nine days after the exper- iment started were below damage threshold levels and did not differ between the treatments. In the May 2011 experiment, the density of sistens crawlers raining down was substantially lower, with 17,000 and 33,000 falling per m2 in the low and high ovisac treatments, respectively. Resulting infestation rates on Carolina hemlock seedlings were extremely low and well below damage threshold levels. Although A. tsugae crawlers were well distributed across the 1 m2 gridded glue sheets placed at the bottom of each cage, hot spots of unusually high crawler density did occur in both experiments. This rain-down technique shows potential for use in an operational tree-breeding program where screening large numbers of hem- lock seedlings for resistance to A. tsugae is required. Keywords: exotic species, host resistance screening, artificial infestation, Tsuga canadensis, Tsuga caroliniana Correspondence: a [email protected], b [email protected], c [email protected] Editor: Brian Aukema was editor of this paper. Received: 27 September 2012 Accepted: 29 March 2013 Published: 1 August 2014 Copyright: This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 14, Number 106 Cite this paper as: Jetton RM, Mayfield III AE, Powers ZL. 2014. Development of a rain down technique to artificially infest hemlocks with the hemlock woolly adelgid, Adelges tsugae. Journal of Insect Science 14(106). Available online: http://www.insectscience.org/14.106 Journal of Insect Science | http://www.insectscience.org 1 Journal of Insect Science: Vol. 14 | Article 106 Jetton et al. Introduction native spruce flora of eastern North America have thus far been immune to A. tsugae infes- The hemlock woolly adelgid, Adelges tsugae tation and successful sexual reproduction has Annand (Hemiptera: Adelgidae), is a serious never been observed in the introduced range. invasive forest pest in the United States, where it poses a significant threat to the long- The first instar nymphs of the progrediens and term sustainability of hemlock (Tsuga spp. sistens generations are called crawlers and are (Pinales: Pinaceae)) ecosystems. The insect the only mobile stage of the insect. After was first described from specimens collected hatching, crawlers move about their natal trees in western North America, where it was ini- or passively disperse to nearby trees in search tially thought to be an exotic pest of western of feeding sites on the pulvinus (needle cush- hemlock (T. heterophylla (Raf.) Sarg.) (An- ion) at the base of hemlock needles, where the nand 1924) but is now known to be native insect remains for the duration of its life (Oten (Havill et al. 2011). The adelgid is exotic in 2011). Adelgids feed by inserting their stylets eastern North America, where it was intro- (piercing/sucking mouthparts) into the plant duced into Virginia in the early 1950s (Havill and extracting stored nutrients from xylem ray et al. 2006), most likely on ornamental nurse- parenchyma cells (Young et al. 1995). Feed- ry stock imported from Japan. The distribution ing causes changes in hemlock cellular of A. tsugae in this region now covers a 19- structures and water potential (Walker-Lane state area, ranging from southern Maine south 2009; Gonda-King et al. 2012) and leads to along the Appalachian Mountain chain to the abortion of reproductive and vegetative northern Georgia and west into Ohio (USDA buds, restriction of new growth, dessication, Forest Service 2011). Throughout much of and needle loss. Severe infestations can kill this range, it has caused widespread decline trees in as few as four years, although some and mortality among populations of eastern trees have been noted to survive infestation hemlock (T. canadensis (L.) Carr.) and Caro- for more than 10 years (McClure et al. 2001). lina hemlock (T. caroliniana Engelm.). The integrated strategy to manage the impact In the eastern United States, A. tsugae has a of A. tsugae on eastern forests includes classi- complex polymorphic life cycle that includes cal biological control, chemical insecticides, two wingless parthenogenetic generations and silvicultural practices, gene conservation, and one winged sexual generation each year host resistance (Onken and Kenna 2008). Of (McClure 1989). The first appearance of each these, biological and chemical control have generation and the timing of life stages varies received the most interest; biological control considerably with latitude (McClure 1989; currently is considered the most promising Gray and Salom 1996; Joseph et al. 2011). tool for long-term adelgid management (Onk- The two parthenogenic generations, called the en and Reardon 2011). Since 2002, however, progrediens and sistens, are present from the general understanding of host‒insect inter- March to June and June to March, respective- actions in the hemlock‒adelgid system has ly. The winged sexual generation is called the improved, and the possibility exists for find- sexuparae and overlaps with the progrediens. ing or breeding A. tsugae-resistant hemlock Upon molting to the adult form, the sexuparae genotypes for reforestation. Initially, both migrate away from hemlock for sexual repro- hemlock species were thought to be universal- duction on spruce (Piceae spp.); however, the ly susceptible to A. tsugae attack (McClure Journal of Insect Science | http://www.insectscience.org 2 Journal of Insect Science: Vol. 14 | Article 106 Jetton et al. 1992), but recent evidence suggests that some needs of larger breeding programs that seek to level of endogenous adelgid tolerance or re- screen among thousands of hemlock geno- sistance may exist within Eastern hemlock types simultaneously. (Caswell et al. 2008) and Carolina hemlock (Jetton et al. 2008, Oten 2011). Attempts to The development of artificial infestation tech- breed the adelgid resistance of T. chinensis niques that are efficient and reliable for (one of the Asian hemlocks) into Carolina screening large numbers of putatively adelgid hemlock through interspecific hybridization resistant genotypes has been identified as a were also successful (Montgomery et al. research priority by the USDA Forest Service 2009). Working Group on Genetics and Host Re- sistance in Hemlock. To address this priority, The screening of tree genotypes for pest re- the preliminary experiments reported here sistance is often accomplished with young were designed to test the utility of a rain-down plants derived from seed or clonally propagat- technique for inducing artificial adelgid infes- ed via rooted stem cuttings or grafted scions. tations on hemlock seedlings en masse. The Upon reaching suitable age or size, these objective of the study was to evaluate the den- plants are artificially infested or inoculated sity and distribution patterns of A. tsugae with the insect or pathogen of concern. Exam- progrediens and sistens crawlers allowed to ples include the selection of pine (Pinus spp.) rain down onto glue sheets and hemlock seed- genotypes for resistance to pitch canker lings. (Fusarium circinatum) and fusiform rust (Cronartium quercuum), and poplar (Populus Materials and Methods spp.) hybrids for resistance to the cottonwood leaf beetle, Chrysomela scripta F. (Foster and A. tsugae source material Anderson 1989; Bingaman and Hart 1992; This study consisted of two separate infesta- Hodge and Dvorak 2007). Within the context tion experiments in the spring of 2011, one of an operational tree breeding program, a key using A. tsugae crawlers of the progrediens
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